J. Nannoplankton Res. 29 (2), 2007, pp.108-128 © 2007 International Nannoplankton Association
ISSN 1210-8049 Printed by Cambridge University Press, UK

108

Six new taxa of subarctic Parmales (Chrysophyceae)
Susumu Konno, Ryoko Ohira
Graduate School of Science & Engineering, Yamagata University, 1-4-12 Kojirakawa-machi, Yamagata 990-8560, Japan

Chika Komuro
Department of Earth & Environmental Sciences, Faculty of Science, Yamagata University, 1-4-12 Kojirakawa-machi, Yamagata 990-8560, Japan

Naomi Harada
Institute of Observational Research for Global Change, Japan Agency for Marine-Earth Science & Technology, 2-15 Natsushima-cho, Yokosuka
237-0061, Japan

Richard W. Jordan*
Department of Earth & Environmental Sciences, Faculty of Science, Yamagata University, 1-4-12 Kojirakawa-machi, Yamagata 990-8560, Japan;*sh081@kdw.kj.yamagata-u.ac.jp
Manuscript received 12th April, 2007; revised manuscript accepted 14th November, 2007

Abstract The Parmales are an enigmatic group of marine phytoplankton, with siliceous plates of varying morphology surrounding their 2-5µm-diameter cells. Although rarely found in sediment traps or underlying sediments,
in oceanic surface-waters they often outnumber even the diatoms during the spring months. Since their discovery,
several decades ago, in the Antarctic and subarctic Pacific, the Parmales have been found in temperate and tropicalregions. Despite the publication of several significant taxonomic papers, a number of Parmales remain without formal description. Here we provide descriptions of six new taxa from subarctic waters: Tetraparma catinifera sp. nov.,
T. gracilis sp. nov., Triparma columacea f. convexa f. nov., T. columacea f. fimbriata f. nov., T. laevis f. inornata f.
nov. and T. laevis f. longispina f. nov. In addition, an annotated checklist of all Parmales, including undescribed taxa,
is presented for the first time. Distribution maps of all subarctic taxa are also provided for the North Pacific and its
marginal seas.

Keywords Bering Sea, Chukchi Sea, Japan Sea, Parmales, Sea of Okhotsk, subarctic Pacific, taxonomy

1. Introduction
Probably due to their small size (2-5µm), the Parmales
escaped detection for most of the 20th Century. However,with the advent of the electron microscope, scientists
began to record the presence of ‘siliceous cysts’ in marine
water samples (Iwai & Nishida, 1976; Nishida, 1979,
1986; Booth et al., 1980, 1981, 1982; Silver et al., 1980;
Takahashi et al., 1986) and of incomplete cysts and isolated siliceous plates in zooplankton faecal pellets (Urban etal., 1993), sediment traps (Ohyama & Jordan, 2002,
unpubl. data) and marine sediments (Stradner & Allram,
1982; Franklin & Marchant, 1995; Zielinski, 1997; Thorn,2004). Booth & Marchant (1987) formally identified
these siliceous cysts as belonging to a new order, theParmales, and assigned them to the Class Chrysophyceae.
The previous year, Marchant & McEldowney (1986) had
sectioned several of these siliceous cysts and demonstrated that they were algae. However, their assignment to the
Chrysophyceae remains speculative, since no one has succeeded in culturing them or conducting genetic studies onthem directly. Recently, Lovejoy et al. (2006) analysed
the genetic diversity of Arctic waters, using 18S rRNA,and noticed several novel sequences. One such cluster
was a sister group to the Bolidophyceae, a group closely
related to the diatoms (Guillou et al., 1999). Lovejoy et al.
(2006) speculated that this cluster represented theParmales, although clearly more evidence is needed
before we can be sure. But if it is true, then Mann &Marchant (1989) may have been right to suggest that a
Parmales-like ancestor gave rise to the diatoms, presumably some time after the Permian/Triassic mass extinction
event (Medlin et al., 1997). However, some of the oldest
Parmales fossils are reportedly those in Late Eocene to
Early Oligocene (ODP Leg 188, Hole 1166A,
148.11mbsf) and Middle to latest Miocene (ODP Leg 188,
Hole 1165B, 169.58-291.28mbsf) sediments from Prydz
Bay, Antarctica (Thorn, 2004). Unfortunately, the author
did not provide photographs to support the findings, andso micrographs of Parmales specimens from the Middle
America Trench slope (DSDP Leg 66, Site 490-1-4, 2526cm), thought to be Middle to Late Quaternary (NN20)
in age, are currently the oldest piece of evidence from thefossil record (Stradner & Allram, 1982).
The exclusively marine Parmales are now well known
from polar waters, however, their occasional presence intropical waters (Silver et al., 1980; Kosman et al., 1993;
Bravo-Sierra & Hernández-Becerril, 2003) suggests that
their distribution may be worldwide, albeit in restricted
habitats. Despite this, the ecology of the Parmales is very
poorly known. Several workers had noticed that theParmales remained covered in plates all year round (e.g.
Booth & Marchant, 1987), but Komuro et al. (2005) wereperhaps the first to show this clearly, and to mention thatthey behaved like diatoms, ‘blooming’ in spring, sinking
to the pycnocline/nutricline in the summer. A few years
earlier, Tanimoto et al. (2003) had shown that, where subarctic Pacific waters entered the Bering Sea (i.e. through109

Konno, Ohira, Komuro, Harada, Jordan

the shallow straits of the Aleutian Islands), the Parmales
were present at the surface during the summer, but awayfrom the Aleutian Islands the Parmales were almost
absent from the subarctic surface-waters. The data ofKomuro et al. (2005) clearly explains why some of theearlier workers found them at deeper depths, while others
found them at the surface. Their seasonal dataset also confirms Marchant & McEldowney’s (1986) observation that
the Parmales are not cysts at all, but the vegetative stageof an alga with a presumably high growth-rate (i.e. higher than that reported by Taniguchi et al. (1995) from theirbag experiments) and high silica uptake rate.
Bravo-Sierra & Hernández-Becerril (2003) provided alist of extant Parmales taxa, including three genera, eightspecies, four subspecies and one forma. They proposed
that, in the future, the status of the four subspecies bechanged to forma or variety, since phytoplankton workers
on other algal groups rarely used subspecies in this way that is, when only morphological features are used as separation criteria, and life-cycle or interbreeding information is not known. Although we support this proposal, we
did not encounter any of the four subspecies in this study,
and so hesitate to make the changes here. However, someof the new taxa herein are assigned forma status, in recognition of their suggestion.
It is now known that the Parmales are a significant
component of the phytoplankton community, especially inhigh latitudes, however their enumeration at the species orsubspecific level is currently hampered by an underestimation of their diversity. Until this problem is addressed,
ecological and biogeographic studies will be difficult toundertake with any confidence. Therefore, in this first taxonomic paper, we describe most of the subarctic taxa thatare presently without formal names.
2. Material and methods
Samples were collected on three cruises undertaken atvarious times of the year and in different years: KH99-3(July-August, 1999) of the R/V Hakuho Maru, MR00K01(January, 2000) and MR06-04 (August-September, 2006)of the R/V Mirai. Figure 1 shows the station locations
from which water-samples were collected and specimens
photographed. The Kyodo North Pacific Ocean TimeSeries (KNOT; Kyodo = cooperative in Japanese) station
was established in 1997 as part of the Joint Global Ocean
Flux Study, and was visited on 16th August, 1999 during
KH99-3 and 17th January, 2000 during MR00K01.
During the first visit, several hydrocasts were carried out
over a short time-interval. Samples from two of thesehydrocasts, referred to in the plate captions simply as
Stations 1 and 2, have been analysed for this study.
Samples were also collected from hydrocasts at Stations
16 and 17 during KH99-3, and from Stations 3, 4, 6, 7 and12ex on MR06-04. In addition, two surface-water samples (Stations 23 and 24) were obtained while underway
on MR06-04. Vertical water-samples were acquired onshallow hydrocasts (5-300m) using a Conductivity

Temperature Depth (CTD) rig equipped with a rosette ofwater bottles. Surface-waters were collected either bybucket, or using the onboard continuous sea-water supply.
For each CTD sample, a suite of hydrographic parameters
was normally measured, while for those taken with theonboard sea-water supply, only temperature and salinity
measurements were available from instruments connectedto the continuous flow of sea-water.
Water-samples were filtered, prepared for scanningelectron microscopy (SEM), and photographed as detailedin Konno & Jordan (2006). All filter samples, SEM stubs,
negatives and scanned images used in this study (including those associated with the holotypes) are presently
curated in the Department of Earth & Environmental
Sciences, Faculty of Science, Yamagata University,
Yamagata, Japan.
3. Results
3.1 Terminology
In general, the terminology used in this paper follows thatof Booth et al. (1981) and Booth & Marchant (1987), butalso incorporates the recent findings and recommendations of Konno & Jordan (2007). The latter authors
revealed that the cell-wall structures of Triparma andTetraparma are more closely related than previously
thought, with the same plate configuration; that is, threegirdle plates, three shield plates, one dorsal plate and one
ventral plate. The major differences between the two genera are the shape of the dorsal and girdle plates and thesize of the ventral plate. In Tetraparma, the triradiate dorsal plate is notched, the girdle plates are also triradiate,
and the ventral plate is smaller than the shield plates. In

Figure 1: Map showing locations of the sampling stations used in thepresent study (closed circles), and those (open circles) of Tanimoto et al.
(2003). Map created using M. Weinelt’s ‘Online Map Creation’ site atwww.aquarius.ifm-geomar.deNew taxa of subarctic Parmales

110

Triparma, the triradiate dorsal plate has rounded ends, thegirdle plates are oblong, and the ventral plate is larger
than the shield plates.

1987 Tetraparma pelagica Booth & Marchant: fig.4.
2003 Tetraparma pelagica Booth & Marchant: Tanimoto et al.,pl.3, fig.6.

3.2 Systematic taxonomy
In this section, six new taxa are described from subarctic
waters. To avoid taxonomic complications in the future,
the bibliography for each taxon has been restricted to thesubarctic Pacific and its marginal seas, although it is presumed that specimens occurring in the subarctic North
Atlantic belong to one or more of the taxa below. For
those species which are seemingly bipolar (i.e. morphologically identical), it is presumed that future studies will
discover that they are actually cryptic species, as has beenfound in other plankton groups. An annotated checklist of
the Parmales, complete with dates and authorities, isgiven in the discussion.
The cell and plate dimensions given in the descriptions below are of specimens photographed in the present
study, while biogeographic references to water-samples
with ‘NP’ and ‘B’ notations are from Tanimoto et al.
(2003). Only one paper provides ultrastructural information on the Parmales. Marchant & McEldowney (1986)
showed that the parmalian cell contains a large chloroplast and very little storage material, indicating its photosynthetic tendencies. Furthermore, they showed that it
possesses a chloroplast endoplasmic reticulum, as foundin diatoms and chrysophytes. Thus, it should be noted thatthe inclusion of the Parmales in the Class Chrysophyceae
sensu lato is still tentative, pending detailed ultrastructural and genetic work.

Etymology: Catina (L.) meaning bowl, fero (L.) meaning
to carry, in reference to the bowl-shaped shield plates.

Class CHRYSOPHYCEAE Pascher
Order PARMALES Booth & Marchant emend.
Konno & Jordan
Family TRIPARMACEAE Booth & Marchant emend.
Konno & Jordan
Genus Tetraparma Booth in Booth & Marchant emend.
Konno & Jordan
Cells planktonic, solitary, non-motile, spherical to subspherical. Cells possess eight plates: three shield plates,
three triradiate girdle plates, one triradiate dorsal plateand one circular ventral plate. Plate boundaries distinct.
All plates are slightly to strongly convex in the centralarea, with or without papillae, and are radially veined withveins dichotomously branching and anastomosing
increasingly toward the margin, forming a wide inner ringof elongate areolae. Arms of triradiate dorsal plate with
notched ends. Marine.
Tetraparma catinifera sp. nov.
Pl.1, figs 1-9; Pl.2, figs 1-2
1976 Sp. indet. A Iwai & Nishida: pl.II, fig.1.
1980 Siliceous cyst Booth et al.: fig.1(8).
1981 Cyst VIII Booth et al.: figs 57-59.

Cellula solitaria, sphaerica, 2.7-3.3µm diametro.
Laminae papillis praesentes, circa 6-12/µm. Papillae
praesentes in seribus radiantibus atque vel concentricis.
Laminae processus centralis, sine umbonatae. Laminae
parmae 1.7-2.4µm diametro, margines elevatos. Laminae
triradiatae, brachii 1.3-1.5µm longitudo. Lamina dorsalis
1.0-1.4µm longitudo. Lamina ventralis, circa 1.6µm
diametro. Species planctonica marina, ad 44˚N, 155˚E(Statio KNOT). Holotypus, hic designatus: EM StubKNOTJ0030. Iconotypus: Lamina 1, Figura 2.
Description: Cells solitary, spherical, 2.7-3.3µm in diameter. Papillae usually present, ca.6-12 papillae/µm. Plates
usually with radially arranged papillae between the slitlike areolae, and one or more concentric rows of papillae
along the top of a raised marginal rim. Plates with small
central papilla, but no central mound. Shield plates
ca.1.7-2.4µm in diameter, with a wall-like plate margin.
Triradiate girdle plates, arms ca.1.3-1.5µm long (measured along the dorso-ventral plane, from end to central
papilla). Arms of dorsal plate, ca.1.0-1.4µm long. Ventral
plate ca.1.6µm in diameter. Marine, in plankton at 44˚N,
155˚E (Station KNOT), 17th January, 2000 (30m).
Holotype: EM Stub KNOTJ0030 (specimen in Pl.1,
fig.2).
Note: In Booth et al. (1981), this form was originally distinguished from T. pelagica (as Cysts VIII and IX, respectively), and when the latter species was formallydescribed in Booth & Marchant (1987), the two forms
were again distinguished in the text, but both appeared in
the figure captions under the same name. Apart from anotable size difference, T. catinifera differs from T. pelagica by having wall-like shield-plate margins of variableheight, lacking the triangular spines at the plate centre,
and having a somewhat flattened central area. However,
Booth & Marchant (1987, fig.5) showed an intriguing
specimen (collected in May from surface-waters at
56˚59’N, 141˚27’W) that seemingly possesses a rimmed
shield-plate in addition to typical T. pelagica plates. In thepresent study though, no such specimens have been seen.
T. catinifera specimens show a significant amount of morphological variation, not just in wall height (compare Pl.1,
fig.1 with Pl.2, fig.1), but also in the distribution anddegree of papillation (compare Pl.1, figs 3, 4 with Pl.1,
figs 6, 7), and the number of radiating slits in the central
area (compare Pl.1, fig.5 with Pl.1, fig.7). It should be
noted that the girdle and dorsal plates have less prominent
walls than the shield and ventral plates (Pl.1, fig.2), andthat, due to the curvature of the cell, all of the plates are
actually convex (Pl.1, fig.9). This species has mostly been

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Konno, Ohira, Komuro, Harada, Jordan

Plate 1
Tetraparma catinifera

Cell wall, showing shield (s), girdle(g), dorsal (d) and ventral (v) plates.
N Pacific, KNOT St.1, 100m

Dorsal plate interlocking with three girdle plates. Note plates strongly papillate.
Sea of Okhotsk, St.7, 30m

Specimen with less prominent
papillae. Ventral plate at bottom of photo. Bering Sea,
KH99-3, St.16, 50m

Specimen clearly showing juncture
between girdle and dorsal (arrowed)
plates. N Pacific, KNOT MR00K01,
30m. Holotype

Specimen with clear plate junctures.
Note papillae on inside of plate rims.
Sea of Okhotsk, St.7, 30m

Note small ventral
plate at bottom ofphoto (arrowed). Sea of
Okhotsk, St.6, 30m

Collapsed cell. Sea of Okhotsk, St.6, 30m

Specimen with moreradiating slits, lacking
papillae. N Pacific, KH99-3
St.17, 50m

Collapsed cell. Note curvature ofplates. Sea of Okhotsk, St.7, 30m

New taxa of subarctic Parmales

112

recorded in the subarctic Pacific, but also occurs in the
Gulf of St. Lawrence, Canada (Bérard-Therriault et al.,1999, p.246, pl.113, figs a, e) and in the Chukchi Sea (thisstudy).
Biogeography (this study): Sea of Okhotsk - MR06-04
St.4, MR06-04 St.6, MR06-04 St.7; NW Pacific - St.
KNOT (August, 1999, KH99-3; January, 2000,
MR00K01), KH99-3 St.17; NE Pacific - KH99-3 NP15;
Bering Sea - MR06-04 St.24, KH99-3 St.16; Chukchi
Sea - MR060-04 St.12ex.

clearly belong to Tetraparma, but are in need of further
observations and formal descriptions. T. gracilis has only
been recorded in the subarctic Pacific.
Biogeography (this study): Sea of Okhotsk - MR06-04
St.7; NW Pacific - St. KNOT (August, 1999, KH99-3),
KH99-3 NP30, KH99-3 St.17; Bering Sea - MR06-04
St.23.

Tetraparma gracilis sp. nov.
Pl.2, figs 3-7

1987 Tetraparma pelagica Booth & Marchant: p.248, figs 2(holotype, from 64˚59.8’S, 83˚02’E; January, surfacewater), 3, 5, non fig.4.
1976 Sp. indet. B Iwai & Nishida: pl.II, fig.2.
1976 Sp. indet. C Iwai & Nishida: pl.II, fig.3?
1979 Genus & species indeterminable Nishida: pl.1, fig.4.
1980 Siliceous cyst Booth et al.: fig.1(9).
1981 Cyst IX Booth et al.: figs 60-62, 69, non fig.68.
2003 Tetraparma pelagica Booth & Marchant: Tanimoto et al.,pl.3, fig.5, non figs 6, 7.

1981 Cyst IX Booth et al.: fig.68?
2003 Tetraparma pelagica Booth & Marchant: Tanimoto et al.,pl.3, fig.7.

Etymology: Gracilis (L.) meaning slender, in reference tothe shape of the central process.
Cellula solitaria, sphaerica, 2.3-2.5µm diametro.
Laminae papillis carentes. Processus centralis, 0.7-0.8µm
longitudo, in laminae sine umbonatae. Laminae margines
laevi et leviter elevatiae. Laminae parmae 1.3-1.8µm
diametro. Laminae triradiatae circa 1µm longitudo.
Lamina dorsalis circa 1µm longitudo. Lamina ventralis,
circa 1.2µm diametro. Species planctonica marina, ad
44˚N, 155˚E (Statio KNOT). Holotypus, hic designatus:
EM Stub KNOTA9960. Iconotypus: Lamina 2, Figura 4.
Description: Cells solitary, spherical, 2.3-2.5µm in diameter. Plates without papillae. All plates lack a central
mound, but have a central process 0.7-0.8µm long.
Processes on girdle and dorsal plates slender, those on
shield and ventral plates appear bifurcate. Plates with asmooth, slightly raised margin. Shield plates ca.1.31.8µm in diameter. Triradiate girdle plates ca.1µm long(measured along the dorso-ventral plane, from end to central structure). Arms of dorsal plate, ca.1µm long. Ventral
plate ca.1.2µm in diameter. Marine, in plankton at 44˚N,
155˚E (Station KNOT), 16th August, 1999 (60m).
Holotype: EM Stub KNOTA9960 (specimen in Pl.2,
fig.4).
Note: T. gracilis differs from T. pelagica by seemingly
lacking papillae and having a long central process. A similar specimen was illustrated by Iwai & Nishida (1976,
pl.II, fig.7), but the central process on at least some of theplates was cruciate. The processes on the shield and ventral plates of some of our specimens appear bifurcate,whilst those on the girdle and dorsal plates are slender(Pl.2, fig.4). Furthermore, in broken specimens, the central process cross-section appears circular (Pl.2, figs 6, 7),
whilst that of T. pelagica (subarctic forms) is elongate(Pl.3, figs 12, 13). Another specimen featured by Nishida(1986, pl.1, fig.4), from the Southern Ocean, has anangled short spine on each plate. Both of these forms

Tetraparma pelagica Booth & MarchantPl.3, figs 1-13

Description: Cells 1.9-2.5µm in diameter. All plates withor without papillae, ca.12-15 papillae/µm, are radiallyveined, with veins dichotomously branching and anastomosing increasingly toward the margin, forming a wide
inner ring of elongate areolae. Plates with radiallyarranged papillae between the slit-like areolae, and twoconcentric rows of papillae on top of a low marginal rim.
Small triangular spine covered by papillae usually present
at plate centre. Shield plates ca.1.3-1.6µm in diameter.
Triradiate girdle plates ca.0.8-1.3µm long (measured
along the dorso-ventral plane from end to central structure). Arms of dorsal plate, ca.0.8µm long. Ventral plateca.1.1-1.8µm in diameter. Marine.
Note: T. pelagica was originally described from theAntarctic (Booth & Marchant, 1987), but smaller, seemingly indistinguishable, cells are also found in the subarctic Pacific and Bering Sea (compare with Findlay, 1998,
pl.3, fig.6; Marchant & Scott, 2005, figs 7.5a, b). Our own
unpublished data on Antarctic T. pelagica specimens also
conform to the original dimensions given by Booth &Marchant (1987). A closer examination of specimensfrom both polar regions may reveal them to be two cryptic taxa in the future. In addition, some specimens lacking
plate spines, but possessing the other characteristic features of this species, have been found exclusively in thesubarctic Pacific (e.g. Iwai & Nishida, 1976, pl.II, fig.3,
as Sp. indet. C; Nishida, 1979, pl.1, fig.4, as Genus &species indeterminable, and in text-figs 4 and 6 as Indet.
B) and the Sea of Okhotsk (Pl.3, figs 10, 11). These nonspiny forms are rare, and presumed to represent the same
species as the spiny forms. The large ventral plate(1.8µm) mentioned in the above description was measured in ventral view, and so one assumes that shield plates
larger than 1.6µm were present on the other side of thecell. A disarticulated specimen of T. pelagica from Disko
Bay, Greenland, clearly shows the size difference between

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Konno, Ohira, Komuro, Harada, Jordan

Plate 2
1-2: Tetraparma catinifera; 3-7: Tetraparma gracilis

Note low-walled rims and less prominent papillae.
Bering Sea, St.24, 75m

Cell wall showing shield (s), dorsal (d) andgirdle (g) plates. N Pacific, KH99-3 St.17,50m

Note dorsal/girdle plate juncture(arrowed). Bering Sea, St.23,
10m

Note low-walled rims and plates with moreradiating slits. N Pacific, KNOT St.1, 100m

Cell wall showing all the plate types, including the ventral
plate (v). N Pacific, KNOT St.1, 60m. Holotype

Note strong curvature of shieldplate in upper left corner. Sea of
Okhotsk, St.7, 30m

Note circular hole left by broken
spine, and ventral plate on lefthand side. Sea of Okhotsk, St.7,30m

New taxa of subarctic Parmales

Plate 3Tetraparma pelagica

Cell wall, showing shield (s),girdle (g), dorsal (d) and ventral(v) plates. Bering Sea, St.24, 0m

Specimen clearly showing dorsal
plates (centre). N Pacific, KH99-3
St.17, 50m

Bering Sea, St.23, 0m

Bering Sea, St.23, 30m

Bering Sea, St.24, 0m

N Pacific, KH99-3 St.17,50m

Note greater degree ofornamentation. Sea of
Okhotsk, St.4, 30m

Note plates seemingly
lack spines. Sea of
Okhotsk, St.7, 30m

Note shape of aperture left bybroken spine on girdle plate, andsmall ventral plate on left. Sea of
Okhotsk, St.7, 30m

Bering Sea, St.23, 10m

Sea of Okhotsk, St.7,30m

Note plates seemingly lackspines. Sea of Okhotsk,
St.7, 30m

Note shape of aperture left by broken
spine on shield plate. Sea of Okhotsk,
St.7, 30m

114

Konno, Ohira, Komuro, Harada, Jordan

115

Plate 4
Triparma columacea f. fimbriata

Cell wall showing girdle (g), shield (s) anddorsal (d) plates (ventral plate not visible). N
Pacific, KNOT St.1, 125m. Holotype

Ventral view showing ventral plate (v). Bering Sea, St.24, 10m

Cell wall with one shield-plate missing. Note ventral platein bottom right corner. Bering Sea, St.23, 30m

Ventral view. Note upper girdle
plates appear interlocked. Bering
Sea, St.24, 30m

Ventral view showing ventral plate. Bering Sea,
St.23, 30m

Ventral view showing ventral plate. Girdle view. Bering Sea, St.23, 0m
Bering Sea, St.24, 50m

New taxa of subarctic Parmales

the shield plates and the ventral plate (Kosman et al.,
1993, fig.31).
Biogeography (this study): Sea of Okhotsk - MR06-04
St.4, MR06-04 St.7; NW Pacific - St. KNOT (August,
1999, KH99-3), KH99-3 St.17; Bering Sea - KH99-3
St.B5, MR06-04 St.23, MR06-04 St.24.
Genus Triparma Booth & Marchant emend. Konno &JordanCells planktonic, solitary, non-motile, spherical to subspherical (excluding extensions). Cells possess eightplates: three shield plates, three oblong girdle plates, one
triradiate dorsal plate, and one circular ventral plate. Plateboundaries distinct. Arms of triradiate dorsal plate withrounded or slightly squarish ends. Marine.
Triparma columacea f. convexa f. nov.
Pl.5, figs 1-4
1976 Sp. indet. E Iwai & Nishida: pl.II, fig.5.
1979 Genus & species indeterminable (Indet. A) Nishida: textfigs 4, 6, pl.1, fig.3.
1981 Cyst VI Booth et al.: pp.71, 74, figs 47-49.
2003 Triparma columacea Booth: Tanimoto et al., pl.3, fig.4.

Etymology: Convexa (L.) meaning convex, in reference
to the shape of the shield plates.
Cellula 1.8-3.3µm diametro. Laminae parmae 1.1-1.7µm
diametro, convexae. Laminae oblongae 1.6-2.1µm longitudo, carina fere conspicua et undulata in aspectu cingularis, seriebus unabus areolae elongatae in utraque lateroet perpendicularibus carinam. Lamina dorsalis brachii
1.0-1.5µm longitudo, fere rotundis extremis. Lamina dorsalis cum foramina centrica, carina triradiata, areolae
elongatae in unibus serie perpendicularibus ad quemquemarginem et formantes latera carinam, atque seribus
pluribus ex areolae parvae in extremum utraque bracho.
Lamina ventralis incognita. Planctonica marina, ad
60˚N, 179˚W (Mare Bering). Holotypus, hic designatus:
EM Stub BS062410. Iconotypus: Lamina 5, Figura 1.
Description: Cells 1.8-3.3µm in diameter. All plates withcoarse venation, without ornamentation. Shield plates
ca.1.1-1.7µm in diameter, convex; radially veined, withveins dichotomously branching and anastomosing
increasingly toward the margin forming a wide inner ringof elongate areolae and a narrow outer ring of compact
areolae. Oblong girdle plates ca.1.6-2.1µm long, withkeel more or less pronounced, undulating in girdle view,
with a single row of elongate areolae on each side of thekeel and perpendicular to it. Arms of dorsal plate, ca.1.01.5µm long with slightly rounded ends. Dorsal plate withcentral indistinct hole, triradiate keel with a single row ofelongate areolae perpendicular to each margin forming
the sides of the keel, and with several rows of small areolae at the end of each arm. Ventral plate unknown. Marine,

116

in plankton at 60˚N, 179˚W, Bering Sea (St.24, 10m).
Holotype: EM Stub BS062410 (specimen in Pl.5, fig.1).
Note: Kosman et al. (1993) have already suggested that it
would be reasonable to describe this taxon as a new form
(their p.119). T. columacea f. convexa is clearly different
from the type (and its currently described forms) as it possesses convex shield-plates rather than flattened ones
(Pl.5, fig.1). Also, there appears to be a difference in the
shape of the ends of the dorsal plate (more squarish in f.
convexa, distinctly rounded in the type: Pl.5, fig.2). In theoriginal description of T. columacea, the dorsal plate is
described as having a hole in the middle, and with small
holes at the end of each arm. Although not clear from our
micrographs, it is possible to see these structures in somespecimens. Furthermore, f. convexa appears to have only
one row of areolae either side of the keel (as in f. alata
Marchant and f. fimbriata), while there are two in thetype. Booth & Marchant (1987, p.251) mentioned thatsimilar forms had been seen in the Bay of Bothnia inwaters of 5-6 PSU (H. Thomsen, pers. comm. in Booth &Marchant, 1987). In a later paper, Kosman et al. (1993)
showed photographs of specimens of T. cf. T. columacea
from Denmark and Finland, but this Baltic Sea form,
although possessing convex shield-plates, had a much
more bulky and less spherical cell shape, due to thestrongly-keeled girdle-plates. In our opinion, the twoforms are different, and both clearly deserve to be separated from the type.
Biogeography (this study): Sea of Okhotsk - MR06-04
St.7; NW Pacific - St. KNOT (January, 2000, MR00K01),
KH99-3 NP30; Bering Sea - MR06-04 St.24.
Triparma columacea f. fimbriata f. nov.
Pl.4, figs 1-7
1980 Siliceous cyst Booth et al.: fig.4.
1981 Cyst IV Booth et al.: pp.68, 71, figs 31-38.
1987 Triparma columacea subsp. alata Marchant in Booth &Marchant: p.251, figs 17, 18 (only N Pacific form).

Etymology: Fimbria (L.) meaning fringe, in reference todistal extension on the girdle plates.
Cellula 2.3-2.7µm diametro. Laminae parmae non altae,
1.3-1.5µm diametro. Laminae oblongae 2.7-3.0µm longitudo, carina cum fimbri ala fere undulata, una serie areolae elongatae in quoque latere et perpendicularis carinam. Lamina dorsali brachii 1.2-1.5µm longitudo, diluterotundis extremis. Lamina dorsalis cum foramine in centro obscurus, carina triradiate, striis elongatis in serie
una perpendicularibus ad quemque marginem et formantibus latera carinum, atque seribus pluribus areolarum
parvarum in extremum utraque bracho. Lamina ventralis
convexa, 2.3-2.7µm diametro, laminae parmae venae
similis, sed annulum angustum exterior areolarum compactarum. Planctonica marina, ad 44˚N, 155˚E, Statio
KNOT. Holotypus, hic designatus: EM Stub

117

Konno, Ohira, Komuro, Harada, Jordan

Plate 5
1-4: Triparma columacea f. convexa; 5-9: Triparma laevis f. laevis

Cell wall showing girdle (g), shield (s)and dorsal (d) plates (ventral plate notvisible). Bering Sea, St.24, 10m.
Holotype

Cell wall with clear radial markings onplates. Bering Sea, St.24, 30m

Dorso-ventral view with ventral plate justvisible (black arrow). Note notch of girdle
plate (white arrow). Sea of Okhotsk, St.7, 30m

Dorsal view with strong keel on
dorsal plate (arrowed). Sea of
Okhotsk, St.7, 30m

Ventral plate (v). Bering
Cell wall showing girdle (g),Sea, St.24, 0m
shield (s) and dorsal (d) plates(ventral plate not visible). Japan
Sea, St.3, 50m

Japan Sea, St.3, 50m

Ventral view. Bering Sea,
St.24, 30m

Ventral view of ventral plate. Bering
Sea, St.23, 0m

New taxa of subarctic Parmales

KNOTA99125. Iconotypus: Lamina 4, Figura 1.
Description: Cells 2.3-2.7µm in diameter. All plates withcoarse venation. Shield plates ca.1.3-1.5µm in diameter,
flattened; radially veined, with veins dichotomously
branching and anastomosing increasingly toward the margin, forming a wide inner ring of elongate areolae. Oblong
girdle-plates ca.2.7-3.0µm long, with a keel bearing an
undulating fringe-like extension, with a single row ofelongate areolae on each side of the keel and perpendicular to it. Arms of dorsal plate ca.1.2-1.5µm long withslightly rounded ends. Dorsal plate with central, indistincthole, triradiate keel, with a single row of elongate areolae
perpendicular to each margin forming the sides of thekeel. Ventral plate somewhat domed, ca.2.3-2.7µm in
diameter, similar venation to shield plates, but with a narrow outer ring of compact areolae. Marine, in plankton at
44˚N, 155˚E, Station KNOT (16th August, 1999, 125m).
Holotype: EM Stub KNOTA99125 (specimen in Pl.4,
fig.1).
Note: T. columacea f. fimbriata differs from the type by
having a projection on the girdle plates, rather than a mere
keel (compare Pl.4, figs 1, 4-7 with Booth & Marchant,
1987, figs 8, 11), and from f. alata by the shape of the projection. In f. fimbriata, the projection extends along theentire length of the girdle-plate (Pl.4, figs 1, 4-7), whereas in f. alata it is located centrally (see Booth &Marchant, 1987, figs 15, 16). Two other forms of T.
columacea are more similar to the type and lack extensions on the girdle plates (see above note on T. columacea
f. convexa).
Biogeography (this study): NW Pacific - St. KNOT
(August, 1999, KH99-3); Bering Sea - MR06-04 St.23,
MR06-04 St.24.
Triparma laevis Booth in Booth & Marchant f. laevis
Pl.5, figs 5-91980 Siliceous cyst Booth et al.: fig.1-1.
1981 Cyst V Booth et al.: p.71, figs 39-46.
1987 Triparma laevis Booth in Booth & Marchant: p.255, figs
31 (holotype, from 56˚45’N, 137˚27’W; May, surfacewater), 32.
2003 Triparma laevis Booth subsp. laevis Tanimoto et al., 2003:
pl.3, fig.2.

Description: Cells 2.4-2.8µm in diameter. Central area ofplates smooth, without papillae or conspicuous areolation.
Shield plates 1.3-1.6µm in diameter, with raised marginal
rim and inverted cone or small arch at centre. Largest of
the girdle plates about 1.8-3.7µm in length, with wing
extending distally 2.0-2.5µm. Each girdle plate has a central spine or two spines, one near each end of the plate.
Spines are buttressed on the dorsal side. Girdle platesinterlock. Arms of dorsal plate, 1.4-1.5µm long, withslightly rounded or somewhat squarish ends and with atriradiate keel, bifurcate at one end. Ventral plate convex,
ca.2.1-2.4µm in diameter, with an incomplete mid-radius
circular ridge and a slightly raised central mound. Marine.

118

Note: This taxon has been recorded in various studies andit always appears to have one girdle wing longer than theothers. However, whether the bifurcation at one end of thedorsal plate keel is always aligned in the same way is notknown as yet. The type, f. laevis, has also been recorded
from the Gulf of St. Lawrence, as T. aff. T. laevis (BérardTherriault et al., 1999, pp.246-247, pl.113f). T. laevis now
has a number of subspecific taxa affiliated to it, whichshare several key characters, but studies have shown thatthe ventral plate ornamentation is a good separation characteristic for this group.
Biogeography (this study): NW Pacific - St. KNOT
(August, 1999, KH99-3; January, 2000, MR00K01); NE
Pacific - NP16; Japan Sea - MR06-04 St.3; Bering Sea- MR06-04 St.23, MR06-04 St.24.
Triparma laevis Booth in Booth & Marchant f. inornata
f. nov.
Pl.6, figs 1-7
1987 “Cell like Triparma laevis” Booth & Marchant: fig.39.

Etymology: Inornatus (L.) meaning unadorned, in reference to the relatively plain girdle plates and lack of plate
spines.
Cellula 2.2-2.7µm diametro. Laminae expolitae, areolae
vel papillis carentes. Laminae parmae 1.4-1.8µm
diametro, convexae, conicae, habens flabella alta centrica. Laminae oblongae 1.9-2.3µm longitudo, carinam ala
similis, spinae carente. Lamina dorsalis brachii 1.51.6µm longitudo, fere rotundis extremis, carina triradiata
non furcata. Lamina ventralis convexa, circa 2.1µm
diametro, flabella alta centrica praesente. Planctonica
marina, ad 60˚N, 179˚W (Mare Bering). Holotypus, hic
designatus: EM Stub BS062430. Iconotypus: Lamina 6,
Figura 1.
Description: Cells 2.2-2.7µm in diameter. All platessmooth, without areolae or papillae. Shield plates ca.1.41.8µm in diameter, convex, conical with a raised flap at
the centre. Oblong girdle plates ca.1.9-2.3µm long, with asingle keel-like wing, lacking spines. Arms of dorsal plateca.1.5-1.6µm long, with slightly rounded ends, triradiatekeel not forked. Ventral plate somewhat domed, ca.2.1µm
in diameter, with raised flaps in the centre. Marine, inplankton at 60˚N, 179˚W, Bering Sea (St.24, 30m).
Holotype: EM Stub BS062430 (specimen in Pl.6, fig.1).
Note: Until now, the other described subspecific taxa of T.
laevis have all possessed some sort of ornamentation; f.
laevis, f. mexicana (Kosman) Hernández-Becerril &Bravo-Sierra and f. longispina have wings, subsp. pinnatilobata Marchant and subsp. ramispina Marchant haveplate spines. However, a somewhat similar form was seenin Antarctica, but the specimen had “a slightly taller triradiate plate keel and heavier wings on the girdle plates”(Booth & Marchant, 1987, p.255, fig.33).

119

Konno, Ohira, Komuro, Harada, Jordan

Plate 6
Triparma laevis f. inornata

Cell wall showing girdle (g), shield (s) anddorsal (d) plates (ventral plate not visible).
Bering Sea, St.24, 30m. Holotype

Ventral view showing ventral plate (v). Sea of
Okhotsk, St.7, 30m

Close-up of dorsal plate. Bering Sea, St.23, 30m

Note plate junctions. Sea of Okhotsk, St.4, 30m

Note plate junctions. Sea of Okhotsk, St.4,30m

Note plate junctions. Bering Sea,
St.23, 10m

Note unusual shield plate. N Pacific,
KH99-3 St.17, 50m

New taxa of subarctic Parmales

Biogeography (this study): Sea of Okhotsk - MR06-04
St.4, MR06-04 St.7; NW Pacific - St. KNOT (August,
1999, KH99-3), KH99-3 St.17; Bering Sea - MR06-04
St.23, MR06-04 St.24.
Triparma laevis Booth in Booth & Marchant f.
longispina f. nov.
Pl.7, figs 1-9
1981 Cyst similar to Cyst V Booth et al.: p.71, figs 66-67.

Etymology: Longus (L.) meaning long, spina (L.) meaning spine, in reference to the long spine on one of the girdle plates.
Cellula 2.8-3.2µm diametro. Laminae expolitae, areolae
vel papillis carente. Laminae parmae 1.6-2.2µm
diametro, convexae, conicae, habens crater centrico.
Laminae oblongae 2.8-3.2µm longitudo, cum ala 0.61.0µm latitudo, margines alam irregularis. Utraque ala
cum spina anterides et longa, 11µm longitudo, extremis
bifurcata. Lamina dorsalis brachii 1.3-2.2µm longitudo,
fere rotundis extremis, carina triradiata et furcata.
Lamina ventralis convexa, 2.5-3.2µm diametro, habens
una flabella centrica et alter flabellae in annulus.
Planctonica marina, ad 59˚N, 179˚W (Mare Bering).
Holotypus, hic designatus: EM Stub BS991650.
Iconotypus: Lamina 7, Figura 3.
Description: Cells 2.8-3.2µm in diameter. All platessmooth, without areolae or papillae. Shield plates ca.1.62.2µm in diameter, convex, conical with a crater at thecentre. Oblong girdle-plates ca.2.8-3.2µm long, with asingle wing 0.6-1.0µm wide, wing margin irregular. Eachwing with a buttressed spine, one of which is very long(up to 11µm), bifurcated at the end. Arms of dorsal plateca.1.3-2.2µm long, with slightly rounded ends, triradiatekeel forked at each end. Ventral plate somewhat domed,
ca.2.5-3.2µm in diameter, with raised flaps in a midradius ring, and one flap in the centre. Marine, in plankton at 59˚N, 179˚W, St.16 (50m).
Holotype: EM Stub BS991650 (specimen in Pl.7, fig.3).
Note: This form was mentioned in Booth & Marchant(1987, p.256) as resembling the Antarctic taxon T. laevis
subsp. pinnatilobata, and in Booth et al. (1981) as being
similar to Cyst V (= T. laevis). Whilst f. longispina isclearly related to T. laevis, it is quite distinct from theother subspecific taxa, with its single, long girdle-spine
and unique ventral-plate ornamentation.
Biogeography (this study): Sea of Okhotsk - MR06-04
St.7; NW Pacific - St. KNOT (August, 1999, KH99-3),
KH99-3 St.17; Bering Sea - KH99-3 St.16, MR06-04
St.23, MR06-04 St.24.
Triparma strigata Booth in Booth & MarchantPl.8, figs 1-10

120

1980 Siliceous cyst Booth et al.: figs 1-7.
1981 Cyst III Booth et al.: p.68, figs 24-30, 65.
1987 Triparma strigata Booth in Booth & Marchant: pp.256,
258, figs 40-42 (holotype = fig.40, from 63˚S, 93˚E;
January, surface-water), ?fig.43
2003 Triparma strigata Booth: Tanimoto et al., pl.3, fig.3.

Description: Cells 2.2-2.8µm in diameter. All platesslightly convex with central area characterised by tubular
processes, 0.2-0.3µm long, sometimes straight or forked,
ca.6-7 processes/µm (Pl.8, figs 4, 6). Plates lack centralarea structure. Shield plates ca.1.3-1.9µm in diameter.
Each girdle plate ca.1.5µm in length, with two straightish
spines, one at each end of the plate, directed at a diverging angle. Spines ca.0.8-6.2µm long and bifurcate at theends. Junction between adjacent girdle plates uncertain.
Arms of dorsal plate ca.1.1-1.5µm long, with squared off
ends. Dorsal plate without a keel, although some of theprocesses are aligned in an identical position to the keels
of other species (Pl.8, figs 1, 7, 9, 10). Other processes on
the dorsal plate are arranged along the plate margin (Pl.8,
figs 7, 9, 10). Ventral plate ca.1.9-2.1µm in diameter.
Plate boundaries sometimes indistinct. Marine.
Note: Although their cell and plate dimensions are smaller, the subarctic Pacific and Bering Sea specimens areotherwise indistinguishable from those illustrated fromthe Antarctic. However, our specimens exhibit a range ofmorphologies. In particular, the length of the girdle
spines; some being very short (Pl.8, fig.3), others much
longer (Pl.8, fig.5). In this study, all of the specimens
appear to bear straight spines, although the spines wereusually broken and so we did not see any spines withbifurcate ends (see Booth et al., 1981, fig.27). In contrast,
specimens from the Gulf of St. Lawrence, Canada, exhibited either straight or twisted spines (Bérard-Therriault etal., 1999, p.246, pl.113b, c, respectively), while Kosman
et al. (1993, figs 33-37, as T. cf. T. strigata) showed an
Antarctic form, which bore shorter girdle-plate spines andshorter plate processes than those specimens attributable
to T. strigata from both polar regions. Furthermore, an
intriguing specimen from the subarctic lacked girdle-plate
spines altogether (fig.42 in Booth & Marchant, 1987),
while a specimen supposedly with mixed Tetraparma/
Triparma features has also been illustrated (fig.30 of
Booth et al., 1981; also shown as fig.43 in Booth &Marchant, 1987). Another area of variability is the central
area. The holotype specimen (Booth & Marchant, 1987)
has a shield plate with a centrally raised structure, as compared to our specimens, which have a more graduallysloped central area (Pl.8, figs 9, 10). The plate junctionsof this form were clearly visible and showed that the endsof the dorsal plate were slightly rounded and fitted into a
correspondingly-shaped notch on the girdle plate(Kosman et al., fig.33). On the other hand, the girdle
plates seemingly abutted with each other (Kosman et al.,fig.35).
Biogeography (this study): Sea of Okhotsk - MR06-04

121

Konno, Ohira, Komuro, Harada, Jordan

Plate 7
Triparma laevis f. longispina

Cell wall showing girdle (g), shield (s) anddorsal (d) plates (ventral plate not visible).
Bering Sea, St.24, 30m

Ventral view showing ventral plate (v).
Bering Sea, St.23, 0m

Note one long, bifurcate girdle spine. Bering Sea,
KH99-3 St.16, 50m. Holotype

Bering Sea, St.23, 30m

Note girdle plate with long spine. N Pacific,
KNOT St.1, 125m

Ventral view. Bering Sea, Ventral plate, girdle view. N
St.23, 0m
Pacific, KNOT St.1, 100m

Bering Sea, St.24, 0m

Ventral plate and junctions with girdle
plates. Bering Sea, St.24, 100m

New taxa of subarctic Parmales

Plate 8Triparma strigata

Cell wall showing girdle (g), shield (s) and dorsal (d)plates (ventral plate not visible). N Pacific, KNOT
St.1, 100m

Ventral view showing ventral plate. Sea of
Okhotsk, St.4, 30m

Ventral view showing ventral plate(v). Bering Sea, St.24, 30m

Cell wall with two long girdle spines (arrowed).
Bering Sea, St.23, 30m

Cell wall with two
Close-up of Fig.5. Note plate
long spines (arrowed). processes are bifurcatedBering Sea, St.24,
30m

Cell wall with one shield plate
missing. Bering Sea, St.24, 0m

Cell wall with one shield
Cell wall with one girdle plate Cell wall with two girdle plates
plate displaced. Note thinness visible (arrowed). Bering Sea, visible (arrowed). Bering Sea,of plates. Bering Sea, St.23, St.24, 50m
St.24, 30m
30m

122

123

Konno, Ohira, Komuro, Harada, Jordan

St.4; NW Pacific - St. KNOT (August, 1999, KH99-3),
KH99-3 NP30; Bering Sea - KH99-3 St.16, MR06-04
St.23, MR06-04 St.24.
Triparma verrucosa Booth in Booth & MarchantPl.9, figs 1-6
1980 Siliceous cyst Booth et al.: fig.1(5).
1981 Cyst I Booth et al.: p.63, figs 3-8, 63-64.
1987 Triparma verrucosa Booth in Booth & Marchant: p.258,
figs 44 (= holotype, from 57˚N, 141˚W; May, surfacewater), 45.

Description: Cells 2.7-3.1µm in diameter. All platesslightly convex with radiating papillae, ca.10-15 papillae/µm (Pl.9, fig.1). On the shield and ventral plates, anumber of papillae are enlarged, some of which form acomplete or incomplete mid-radius ring. The ventral plate
may have another ring near the margin (Pl.9, fig.2).
Circular plates lack central-area structure. Shield plates
ca.1.7-1.8µm in diameter. Each girdle plate 1.7-2.1µm in
length, with two straightish spines, up to ca.7µm long,
one at each end of the plate, directed at a diverging angle.
Narrow wing margin usually straight, occasionally crenulate. Girdle-plate papillae in rows perpendicular to thewings, adjacent plates interlock by alternating rounded
and notched ends. Arms of dorsal plate ca.1.4-1.5µm
long, with squared off ends. Dorsal plate with keel, bothends of keel forked (Pl.9, figs 1, 6). Dorsal plate papillae
in rows perpendicular to plate arms. Ventral plate ca.2.32.8µm in diameter. Marine.
Note: In the original description of this species, Booth &Marchant (1987) mentioned that the spines on the girdle
plates are ca.5µm long, however, those on one of our
specimens shown here are considerably longer (see Pl.9,
fig.3). Furthermore, a specimen from St. KNOT (collected in August, 1999), not shown here, had a girdle spinewith a bifurcate end. However, this specimen photographed in ventral view had a different ventral platemorphology to those shown here. Our observations
appear to be the first record of this species on the westernside of the Pacific and in the Sea of Okhotsk.
Biogeography (this study): Sea of Okhotsk - MR06-04
St.6, MR06-04 St.7; NW Pacific - St. KNOT (August,
1999, KH99-3).
4. Discussion
4.1 New taxa
In this study, a total of 10 taxa were encountered in our
samples, six of which are formally described as new. As aresult, most Parmales in the subarctic Pacific region can
now be identified with a higher degree of confidence.
However, some of these described taxa are in need of further observations (e.g. the appearance of the ventral plateof Triparma columacea f. convexa is currently unknown).
Also, several subarctic taxa previously featured by otherworkers were absent from our samples (e.g. Triparma

columacea f. columacea, T. retinervis and T. retinervis
subsp. crenata). Despite this, we believe that great
progress is being made on the taxonomy of this understudied, yet significant, group of phytoplankton, and weare now in a better position to carry out detailed seasonaland biogeographic studies using our extensive sample
collection. Additional studies are currently being undertaken in the Antarctic, where similar taxonomic uncertainty exists, and so it is hoped that the true diversity of thisgroup may be realised in the near future. In an attempt tohasten this process, an annotated checklist of Parmales
taxa is presented below, which includes undescribed taxa
and those potentially cryptic taxa which are currently considered to have a bipolar distribution.
4.2 Checklist of extant and fossil
ParmalesClass Chrysophyceae Pascher, 19141
Order Parmales Booth & Marchant, 1987 emend. Konno& Jordan, 2007
Family Pentalaminaceae Marchant in Booth & Marchant,
1987 emend. Konno & Jordan, 2007
Pentalamina Marchant in Booth & Marchant, 1987
P. corona Marchant in Booth & Marchant, 19872
Family Triparmaceae Booth & Marchant, 1988 emend.
Konno & Jordan, 20073
Tetraparma Booth in Booth & Marchant, 1987 emend.
Konno & Jordan, 2007
T. catinifera Konno et al., 2007
T. gracilis Konno et al., 2007
T. insecta Bravo-Sierra & Hernández-Becerril, 20034
T. pelagica Booth & Marchant, 19875
Sp. indet. G sensu Iwai & Nishida, 19766
Siliceous microorganism sensu Nishida, 19867
Siliceous cyst sensu Silver et al., 1980, fig.38
Enigmatic siliceous cyst sensu Stradner & Allram, 19829
Triparma Booth & Marchant, 1987 emend. Konno &Jordan, 2007
T. columacea subsp. alata Marchant in Booth &Marchant, 1987
T. columacea Booth in Booth & Marchant, 1987 f.
columacea
T. columacea f. convexa Konno et al., 2007
T. columacea f. fimbriata Konno et al., 2007
T. cf. T. columacea sensu Kosman et al. 1993, figs 17-2310
T. laevis f. inornata Konno et al., 2007
T. laevis Booth in Booth & Marchant, 1987 f. laevis
T. laevis subsp. laevis (Antarctic form) sensu Booth &Marchant, 198711
T. laevis f. longispina Konno et al., 2007
T. laevis f. mexicana (Kosman in Kosman et al., 1993)
Hernández-Becerril in Bravo-Sierra & HernándezBecerril, 200312
T. laevis subsp. pinnatilobata Marchant in Booth &Marchant, 1987

New taxa of subarctic Parmales

Plate 9Triparma verrucosa

Cell wall showing girdle (g), shield (s) and dorsal (d)plates (ventral plate not visible). Sea of Okhotsk, St.6,30m

Ventral view showing ventral plate (v). Sea of Okhotsk, St.6,
30m

Specimen with long girdle spines. N Pacific, KNOT
St.2, 60/70m

Ventral view with ventral plate inside collapsed cell. Note girdleplate junctures (arrowed). Sea of Okhotsk, St.7, 30m

Collapsed cell in ventral view. Sea of Okhotsk,
St.7, 30m

Cell in dorsal view. Note notch in girdle plate (arrowed). Sea of
Okhotsk, St.7, 30m

124

125

Konno, Ohira, Komuro, Harada, Jordan

T. laevis subsp. ramispina Marchant in Booth &Marchant, 1987
T. retinervis subsp. crenata Booth in Booth & Marchant,
1987
T. retinervis Booth in Booth & Marchant, 1987 subsp.
retinervis
T. strigata Booth in Booth & Marchant, 198713
T. verrucosa Booth in Booth & Marchant, 1987
Sp. indet. F sensu Iwai & Nishida, 197614
Spined forms of Triparma spp. sensu Kosman et al.,199315
Cysts of uncertain affiliation
Unidentified species of Parmales sensu Tanimoto et al.,200316
Cyst 3D sensu Takahashi et al., 198617
Cyst 3A sensu Takahashi et al., 198618
Cyst 3B sensu Takahashi et al., 198619
Cyst 4C sensu Takahashi et al., 198620
Taxonomic notes
1 Pascher (1914) was the first person to erect the Class
Chrysophyceae, and since then it has been emended manytimes as groups of taxa were removed or added (e.g.
Christensen, 1962; Hibberd, 1976). Here, we follow the
stance taken by Kristiansen & Preisig (2001), in which
Chrysophyceae sensu lato is retained, and in which theOrder Parmales can be included.
2 Previously recorded as ‘Cyst 4A’ by Takahashi et al.
(1986, figs 17, 18), and also illustrated by Silver et al.
(1980, figs 1C, E) and Buck & Garrison (1983, fig.35)
from Antarctic waters.
3 Replaced original family name, Octolaminaceae Booth& Marchant, 1987, which was invalid since it was notbased on a generic name (ICBN Art.18.17: Voss et al.,1983 Edition).
4 Only reported from the Gulf of Tehuantepec, Mexican
Pacific (Bravo-Sierra & Hernández-Becerril, 2003).
5 Presently considered to be bipolar, however, some variation has been noted (Booth & Marchant, 1987). The
holotype from the Antarctic has spines, as do most specimens in the subarctic Pacific (e.g. Iwai & Nishida, 1976,
pl.II, fig.2, as Sp. indet. B; Tanimoto et al., 2003, pl.3,
fig.5), while some specimens in the subarctic Pacific lackthem (e.g. Iwai & Nishida, 1976, pl.II, fig.3, as Sp. indet.
C; Nishida, 1979, pl.1, fig.4, as Genus & species indeterminable and Indet. B). Future studies may show that specimens from the subarctic belong to a cryptic species, but
in the meantime the name T. pelagica should be used.
6 This form, illustrated by Iwai & Nishida (1976, pl.II,
fig.7), resembles T. gracilis, but differs in possessing
cross-shaped central spines. Clearly, more specimens areneeded before this taxon can be formally described.
7 This form, illustrated by Nishida (1986, pl.1, fig.4) andKonno & Jordan (2007, fig.4), also resembles T. gracilis,
but differs in having shorter, more pointed central spines.
8 This form slightly resembles T. insecta, but has highly-

domed circular shields, with a small papilla on top. It wasrecorded in a towed net sample from 2800m water-depth
in the eastern equatorial Pacific (Silver et al. 1980, fig.3).
It appears to have a cell wall composed of shield and triradiate plates, and so belongs in the genus Tetraparma.
9 Specimens of an ‘enigmatic siliceous cyst’ found in
Middle-Late Quaternary sediments off Mexico (Stradner& Allram, pl.1, figs 1-4) resemble the form found by
Silver et al. (1980), but differ in having a raised structurewith straighter edges, on top of which there is a papillate
hump. This form appears to have a cell wall composed ofshield and triradiate plates, and so should be placed in thegenus Tetraparma.
10 The holotype specimen of T. columacea, and otherspecimens from the type material (subarctic Pacific), haveflattened shield plates, while those from Finland andDenmark (Kosman et al., 1993, figs 17-23) have convex
centres. Specimens of the latter differ from T. columacea
f. convexa by having a much more bulky and less spherical cell-shape due to strongly keeled girdle-plates.
11 Booth & Marchant (1987, fig.33) showed a specimen ofT. laevis subsp. laevis from the Antarctic. However, its
girdle plates lacked the extended wings which characterise the holotype. Konno & Jordan (2007, figs 5, 6)
showed two more specimens from the Antarctic, as T. laevis f., one of which possessed a different ventral plate thanthat of T. laevis f. laevis. This taxon (or taxa) needs to be
examined more thoroughly.
12 Originally described from the Sea of Cortez as T. laevis
subsp. mexicana Kosman (Kosman et al., 1993, figs 110).
13 T. strigata was originally described from the Antarctic(Booth & Marchant, 1987, figs 40, 41), although similar
specimens had previously been recorded from the subarctic (Booth et al., 1980, 1981). A spineless form was also
found in the subarctic (Booth & Marchant, 1987, fig.42),
and a specimen with twisted spines was illustrated fromthe Gulf of St. Lawrence (Bérard-Therriault et al., 1999,
pl.113c), but these are not considered here to be separate
taxa. Also, it is not possible to separate the Northern andSouthern Hemisphere specimens on purely morphologicalgrounds, and so for the time being the name T. strigata
should be used.
14 Iwai & Nishida (1976, pl.II, fig.6, as Sp. indet. F) illustrated a specimen from 50˚20’N, 178˚56’E that had anelongate central process on each shield plate.
15 Cyst II of Booth et al. (1981, figs 9-16, 21) included
specimens with and without long girdle spines. Later,
Booth & Marchant (1987, figs 23, 24, 27, 28) described T.
retinervis subsp. crenata, based on the spineless forms,
but noted that a “form of Triparma retinervis subsp. crenata with spines” also existed (p.253, fig.29). Kosman etal. (1993, figs 24-28) illustrated spined forms of Triparma
spp. from California and Antarctica, while isolated scales,
called ‘pulvinate siliceous structures’, previously featured
by Norris (1971, pl.2, figs 10-13; pl.3, fig.14) from theIndian Ocean, may belong to a similar taxon.

New taxa of subarctic Parmales
16 A spiny form first illustrated by Tanimoto et al. (2003,

pl.3, fig.1) from NP38 (east of the Tsugaru Strait), wasfound in the surface-waters of the Soya Strait (MR06-04
St.3) in this study. Although Tanimoto et al. (2003, p.102,
pl.3, fig.1) referred to it as an “unidentified species ofParmales”, it has overlapping monomorphic scales, andso lacks the dimorphic or polymorphic interlocking platescharacteristic of all Parmales genera.
17 Cyst 3D, from the Kita-no-seto Strait near Syowa
Station in Antarctica (Takahashi et al., 1986, pl.3, fig.14),
appears to be related to the Parmales (Booth & Marchant,
1987). In particular, the bottom left specimen has lost oneof its ‘shield’ plates, and the resulting view of the flange
looks similar to that seen in specimens of Parmales.
18, 19 Cyst 3A and Cyst 3B, from the Kita-no-seto Strait
near Syowa Station in Antarctica (Takahashi et al., 1986,
pl.2, figs 9, 10 and 11, 12 respectively) may or may not be
related to the Parmales (Booth & Marchant, 1987). This
uncertainty is due to the fact that the ‘cysts’ comprise twohemispherical parts, supposedly with flattened bottoms(only their pl.2, fig.12 shows what appears to be a dimpled apex), and a narrow flange. Cyst 3B also has long,tapering spines associated with it. Clearly, more specimens are needed to elucidate the true nature of these‘cysts’, but all the Parmales known so far have five oreight plates, with two to four plate types that are morphologically different.
20 Booth & Marchant (1987) believed that Cyst 4C of
Takahashi et al. (1986, figs 22, 23) belongs to an undescribed species of Pentalamina, but since then no one has
reinvestigated this form. The cell wall appears to be composed of three round plates and two triradiate plates.
4.3 Biogeographic distribution ofsubarctic taxa
Photographic data from the subarctic Atlantic is rather
sparse, limited to isolated sampling points in the Gulf of
St. Lawrence (Bérard-Therriault et al., 1999), Gulf ofBothnia in Finland (Thomsen, 1986; Kosman et al.,1993), Great Belt in Denmark, and Disko Bay inGreenland (Kosman et al., 1993). Consequently, biogeographic maps encompassing only the subarctic Pacificand its marginal seas were compiled for each taxon, using
data from the literature as well as that generated during
this study (Figure 2). Although this dataset is also limited,
it is clear that many of the taxa are distributed across thesubarctic zone, whilst others appear poorly distributed
due to their rarity and the paucity of samples analysed forParmales.
5. Conclusions
In this study, ten taxa of Parmales have been recorded andillustrated from the subarctic Pacific and its marginalseas, including the descriptions of six new taxa. As aresult of these additions, a checklist of all Parmales has
been presented, with the aim of providing some stimulus
for further taxonomic research. Using the data collected126

thus far, biogeographic maps of each taxon have been
compiled for the study area. Perhaps from now on,
detailed ecological studies can be carried out with greatertaxonomic certainty.
Acknowledgements
The authors would like to thank the captains, crews and scientists involved in the various cruises mentioned above for their
help and cooperation in acquiring the CTD data and water-samples used in this paper. We are also grateful to former students,
Chieko Aizawa, Yumiko Iwasaki and Maiko Tanimoto whose
thesis datasets, negative collections and SEM stubs have provided us with additional information. This research was partly supported by a Grants-in-Aid for Scientific Research (No.
13440152) awarded to RWJ and Kozo Takahashi by theJapanese Society for the Promotion of Science (JSPS). Finally,
we would like to thank the reviewers, Harvey Marchant andDavid Hernández-Becerril, and the Editor, Jackie Lees, for theirvaluable comments and suggestions on our manuscript, especially David Hernández-Becerril who greatly improved theLatin diagnoses.

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